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          细胞能量的转换
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            <p>细胞生物学 第六章 线粒体和叶绿体</p>
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<hr>
<h2 id="线粒体"><a href="#线粒体" class="headerlink" title="线粒体"></a>线粒体</h2><p>线粒体(“动力工厂”)将有机化学能转换成了ATP，</p>
<p>发现过程：</p>
<ul>
<li>1890年，德国生物学家 Altmann动物细胞颗粒结构“生命小体”</li>
<li>1897年， Benda命名之“线粒体”</li>
<li>1904年， Meves在植物细胞发现“线粒体”</li>
</ul>
<h3 id="线粒体的形态"><a href="#线粒体的形态" class="headerlink" title="线粒体的形态"></a>线粒体的形态</h3><p>借助光学显微镜， 早期的科学家观察到的线粒体，正如命名时的取意， 呈颗粒或短线状。</p>
<p>【大小】：直径为0.3~1.0μm，长度为1.5~3.0μm</p>
<p>但在许多动、植物细胞或特定细胞周期时相中， 线粒体的大小和形态可能随着细胞生命活动的变化而呈现很大的变化。</p>
<blockquote>
<p>比如， 人成纤维细胞线粒体可长达40 μm , 植物分生组织细胞中会出现环核的片层状线粒体等。</p>
</blockquote>
<p>【数量】：同一类型细胞中线粒体数目相对稳定；不同类型细胞中线粒体数目相差较大</p>
<blockquote>
<p>衣藻和红藻等低等的真核细胞每个细胞只含有一个线粒体， 而高等动物细胞内含有数百到数千个线粒体， 说明细胞中线粒体的数目受到物种遗传信息的调控。</p>
<p>在同一种高等动植物体内， 细胞内线粒体数目与细胞类型相关， 说明细胞内线粒体的数目随着细胞分化而变化。</p>
</blockquote>
<p>【分布】：一般不均匀，能量需求集中的区域线粒体分布密集。</p>
<h3 id="线粒体的超微结构"><a href="#线粒体的超微结构" class="headerlink" title="线粒体的超微结构"></a>线粒体的超微结构</h3><p><img src="http://img.whl123456.top/image/%E7%BA%BF%E7%B2%92%E4%BD%93%E8%B6%85%E5%BE%AE%E7%BB%93%E6%9E%84%E7%9A%84%E6%A8%A1%E5%BC%8F%E5%9B%BE.png" alt="线粒体超微结构的模式图"></p>
<h4 id="外膜"><a href="#外膜" class="headerlink" title="外膜"></a>外膜</h4><p>单位膜，光滑，厚约6nm</p>
<p>蛋白质/脂类=1：1</p>
<p>外膜上分布有<strong>孔蛋白</strong>（porin）构成的桶状通道，直径为2~3 nm，可根据细胞的状态可逆性地开闭。</p>
<p>当孔蛋白通道完全打开时，可以通过相对分子质量高达5×10<sup>3</sup> 的分子。ATP、NAD、辅酶A 等相对分子质最小于1×10<sup>3</sup> 的物质均可自由通过外膜。由于外膜的<strong>通透性很高</strong>， 膜间隙中的离子环境几乎与胞质相同。</p>
<p><img src="http://img.whl123456.top/image/%E5%AD%94%E8%9B%8B%E7%99%BD.png" alt="孔蛋白"></p>
<p>【外膜功能】：参与膜磷脂的合成，对将在线粒体基质中彻底氧化的物质进行先行初步分解。</p>
<p>外膜上还分布有一些特殊的酶类， 如参与肾上腺素氧化、色氨酸降解、脂肪酸链延长的酶等， 表明外膜不仅参与膜磷脂的合成， 还可对将在线粒体基质中彻底氧化的物质进行先行初步分解。外膜的<strong>标志酶是单胺氧化酶</strong>（monoamine oxidase） 。</p>
<h4 id="内膜"><a href="#内膜" class="headerlink" title="内膜"></a>内膜</h4><p>单位膜；封闭系统膜；厚6-8nm</p>
<p>相对外膜而言， 内膜有很高的蛋白质／ 脂质比（ 质量比≥3 : 1 ) 。</p>
<p>缺乏胆固醇，富含心磷脂（约占磷脂含最的20%），通透性低。</p>
<blockquote>
<p>这种组成决定了内膜的不透性( impermeability） ， 从而限制了所有分子和离子的自由通过， 是质子电化学梯度的建立及ATP合成所必需的。</p>
</blockquote>
<p>特异载体蛋白</p>
<p>ATP合酶</p>
<p><img src="http://img.whl123456.top/image/ATP%E5%90%88%E9%85%B6.png" alt="ATP合酶"></p>
<p>内膜表面不光滑</p>
<p>电子传递链</p>
<p><img src="http://img.whl123456.top/image/%E7%94%B5%E5%AD%90%E4%BC%A0%E9%80%92%E9%93%BE.png" alt="电子传递链"></p>
<p>线粒体内膜向内延伸形成嵴， 大大增加了内膜的表面积。</p>
<blockquote>
<p>据测算， 肝细胞线粒体内膜的表面积相当千外膜的5倍， 细胞质膜的17倍</p>
</blockquote>
<p>嵴的形状、数量和排列与细胞种类及生理状况密切相关。通常情况下， 能最需求较多的细胞中线粒体嵴的数量也较多</p>
<blockquote>
<p>心肌和骨骼肌线粒体嵴的数最相当于肝细胞线粒体峭的3倍等。</p>
<p>这种差异可能反映了不同组织细胞对ATP的需求。</p>
</blockquote>
<p>【内膜功能】：电子传递和氧化磷酸化的主要部位</p>
<h4 id="膜间隙"><a href="#膜间隙" class="headerlink" title="膜间隙"></a>膜间隙</h4><p>膜间隙的宽度通常维持在6~8nm。</p>
<p>当细胞活跃呼吸时， 膜间隙可显著扩大。</p>
<p>膜间隙内的液态介质含有可溶性的酶、底物和辅助因子。</p>
<p>腺甘酸激酶是膜间隙的标志酶， 其<strong>功能</strong>为催化ATP分子末端磷酸基团转移到AMP, 生成ADP</p>
<h4 id="线粒体基质"><a href="#线粒体基质" class="headerlink" title="线粒体基质"></a>线粒体基质</h4><p>线粒体基质为富含可溶性蛋白质的胶状物质， 具有特定的pH和渗透压。</p>
<p>催化线粒体重要生化反应</p>
<ul>
<li>三羧酸循环</li>
<li>脂肪酸氧化</li>
<li>氨基酸降解</li>
<li>丙酮酸降解</li>
</ul>
<p>基质中还含有DNA、RNA、核糖体以及转录、翻译所必需的重要分子</p>
<p>【功能】：三羧酸循环；脂肪酸氧化；线粒体DNA复制转录翻译</p>
<hr>
<p>线粒体的化学成分</p>
<ul>
<li>蛋白质65-70%</li>
<li>脂类25-30%</li>
<li>DNA、RNA</li>
</ul>
<div class="tabs" id="小标签"><ul class="nav-tabs"><li class="tab active"><a href="#小标签-1">外膜</a></li><li class="tab"><a href="#小标签-2">膜间隙</a></li><li class="tab"><a href="#小标签-3">内膜</a></li><li class="tab"><a href="#小标签-4">基质</a></li></ul><div class="tab-content"><div class="tab-pane active" id="小标签-1"><ul>
<li>细胞色素b<sub>5</sub></li>
<li>NADH细胞色素c还原酶</li>
<li><strong>单胺氧化酶</strong></li>
<li>脂酰辅酶A合成酶</li>
<li>磷酸甘油酰基转移酶</li>
<li>孔蛋白</li>
</ul></div><div class="tab-pane" id="小标签-2"><ul>
<li><strong>腺苷酸激酶</strong></li>
<li>核苷酸激酶</li>
<li>磷酸激酶</li>
<li>单磷酸激酶</li>
</ul></div><div class="tab-pane" id="小标签-3"><ul>
<li>NADH脱氢酶</li>
<li>琥珀酸脱氢酶</li>
<li><strong>细胞色素氧化酶</strong></li>
<li>细胞色素C</li>
<li>ATP合成酶（F<sub>0</sub>F<sub>1</sub>复合物）</li>
<li>运输蛋白</li>
</ul></div><div class="tab-pane" id="小标签-4"><ul>
<li>丙酮酸脱氢酶</li>
<li>脂肪酸β氧化化酶</li>
<li>三羧酸循环酶系</li>
<li><strong>苹果酸脱氢酶</strong></li>
<li>DNA聚合酶</li>
<li>RNA聚合酶</li>
<li>核糖体</li>
<li>转移RNAs</li>
</ul></div></div></div>

<h3 id="氧化磷酸化"><a href="#氧化磷酸化" class="headerlink" title="氧化磷酸化"></a>氧化磷酸化</h3><p>在生物分子氧化过程中，伴有ADP磷酸化生成ATP的作用，即氧化与磷酸化相偶联</p>
<p>磷酸化作用</p>
<p>动物两种方式：底物水平磷酸化；电子传递磷酸化（产生95%的ATP）</p>
<p>植物三种方式：底物水平磷酸化；电子传递磷酸化；光合磷酸化</p>
<p>氧化磷酸化通常特指线粒体中的电子传递磷酸化作用，本质是电子的转移</p>
<h4 id="电子载体与电子传递链"><a href="#电子载体与电子传递链" class="headerlink" title="电子载体与电子传递链"></a>电子载体与电子传递链</h4><p>电子载体</p>
<ul>
<li>NAD<sup>+</sup>、NADP<sup>+</sup> </li>
<li>FMN、FAD</li>
<li>铁硫中心（铁-硫簇）</li>
<li>辅酶Q（泛醌）</li>
<li>细胞色素类a a<sub>3</sub> b c c<sub>1</sub></li>
<li>Cu</li>
</ul>
<hr>
<p>通用电子载体</p>
<p>NAD<sup>+</sup>：烟酰胺腺嘌呤二核苷酸</p>
<p>NADP<sup>+</sup> ：咽酰胺腺嘌呤二核苷酸磷酸</p>
<hr>
<p>黄素蛋白类电子受体</p>
<p>FMN：黄素单核苷酸</p>
<p>FAD：黄素腺嘌呤二核苷酸</p>
<hr>
<p>铁硫中心（铁-硫簇）</p>
<p><img src="../../../../../Pictures/blogimg/%E9%93%81%E7%A1%AB%E4%B8%AD%E5%BF%83.png" alt="铁硫中心"></p>
<hr>
<p>辅酶Q（泛醌）</p>
<p>线粒体内膜上的脂溶性小分子，可在脂双层中自由移动</p>
<hr>
<p>细胞色素类</p>
<p>一类以卟啉为辅基的电子传递蛋白</p>
<p>Cyt a a<sub>3</sub> b c c<sub>1</sub></p>
<hr>
<p>电子呼吸链</p>
<ul>
<li><p>NADH呼吸链</p>
</li>
<li><p>FADH2呼吸链</p>
</li>
</ul>
<hr>

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